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Fig. 3.1 Co-localization of kisspeptin with other neuropeptides/transmitters. Examples of the
dual-labeling of ARC kisspeptin cells ( top row ) with dynorphin (DYN) and neurokinin B (NKB)
in sheep, rats, and monkeys, and of RP3V kisspeptin cells ( bottom row ) with tyrosine hydroxylase
(TH, a marker for dopamine) and met-enkephalin (mENK) in mice. Dual-labeled cells appear yel-
low in ( a - d ), and white in ( e ) (indicated by arrows ). Arrow in ( b ) shows an example of a close
contact between a DYN/NKB terminal and a DYN/NKB (KNDy) cell in the ARC. Images are
modifi ed with permission from [ 10 ] ( a ); [ 105 ] ( b ); [ 40 ] ( c ); [ 64 ] ( d ); and [ 67 ] ( e ). Scale bars in
( a ) 20 μm, ( b ) 10 μm, ( c ) 20 μm, ( d ) 40 μm, and ( e ) 20 μm. ( a ) Modifi ed from Goodman RL,
Lehman MN, Smith JT, Coolen LM, De Oliveira CVR, Jafarzadehshirazi MR et al. Kisspeptin
neurons in the arcuate nucleus of the ewe express both dynorphin A and neurokinin B.
Endocrinology 2007; 148(12):5752-5760 (with permission from The Endocrine Society).
( b ) Modifi ed from Burke MC, Letts PA, Krajewski SJ, Rance NE. Coexpression of dynorphin and
neurokinin B immunoreactivity in the rat hypothalamus: morphologic evidence of interrelated
function within the arcuate nucleus. Journal of Comparative Neurology 2006; 498(5):712-726
(with permission from John Wiley & Sons). ( c ) Modifi ed from Ramaswamy S, Seminara SB,
Ali B, Ciofi P, Amin NA, Plant TM. Neurokinin B stimulates GnRH release in the male monkey
(Macaca mulatta) and is co-localized with kisspeptin in the arcuate nucleus. Endocrinology 2010;
151(9):4494-4503 (with permission from The Endocrine Society). ( d ) Modifi ed from Clarkson J,
Herbison AE. Dual phenotype kisspeptin-dopamine neurones of the rostral periventricular area
of the third ventricle project to gonadotropin-releasing hormone neurones. Journal of
Neuroendocrinology 2011; 23(4):293-301 (with permission from John Wiley & Sons).
( e ) Modifi ed from Porteous R, Petersen SL, Yeo SH, Bhattarai JP, Ciofi P, de Tassigny XD et al.
Kisspeptin neurons co-express met-enkephalin and galanin in the rostral periventricular region of
the female mouse hypothalamus. Journal of Comparative Neurology 2011; 519(17):3456-3469
(with permission from John Wiley & Sons)
co-localization is uniformly high among female mammals and male rodents and
ruminants (Table 3.3 ), it is slightly lower in male monkeys [ 40 ] and humans [ 44 ],
even in the absence of gonadal steroids that inhibit kisspeptin expression in the
ARC. In addition, perturbations in the organizational effects of gonadal steroids
during development can lead to an altered ratio of KNDy peptides within the adult
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